grounding language on neurobiology · grounding language on neurobiology angela d. friederici ....
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Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Max Planck Institute for Human Cognitive and Brain Sciences Leipzig, Germany
Grounding language on neurobiology
Angela D. Friederici
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Fitch.ppt
courtesy of T. Fitch
The language faculty is human specific. What is the neurobiological basis of
this faculty?
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Friederici, Physiological Reviews, 2011
Fachbeirat_2012_NP.ppt
LH RH
Language in the human brain
Language is realized in the human brain as computations in specialized cortical areas that are tightly coupled functionally and structurally to form a large scale network for language processing. It receives its specificity due to syntax, a system of rules that permits the combination, organization and permutation of words in meta-structures, i.e. sentences.
Structure Structure Function Function
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Pattern for non-embedded sentences
Pattern for embedded sentences
Basic neurophysiological principles
Amsterdam, 29.08.2014 >> Bilder >> Meyer.ppt
Although human specific, language is based on common neurophysiological principles. Individual neurons and neuronal assemblies are the basis of neuronal activity which for language usually can only be measured as spatiotemporal pattern at the cortical level (except for intracranial recordings).
Source: Meyer, Grigutsch, Schmuck, Gaston & Friederici, Poster presented at CNS, 2014
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
160 ms
550 ms
700 ms
ELAN
N400
P600 -3.0
3.0 µV
-2.0
2.0 µV
-3.0
3.0 µV
access to syntactic category info & phrase structure building
access to lexical-semantic info &
thematic role assignment
Tim
e
integration
Source: 3-Phases Model by Friederici, Brain and Language, 1995; Trends in Cognitive Sciences, 2002
EEG:Neurochronometry of language comprehension
EEG data have allowed to identify specific ERP components reflecting different aspects and levels during on-line sentence comprehension.
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Localization of the ELAN effect
left hemisphere right hemisphere
#1
#2
#3
#4
#5
158 ms
142 ms
133 ms
143 ms
139 ms
#3
133 ms left hemisphere right hemisphere
Source: Friederici et al., Human Brain Mapping, 2000
Phase 1: MEG localization of the ELAN
Using MEG the ELAN representing initial phrase structure processes could be localized in the anterior temporal and inferior frontal cortex.
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Phase 2: The N400 allows to differentiate different aspects
Source: Hagoort, Hald, Bastiaansen & Petersson, Science, 2004
Semantic N400-effect World knowledge N400-effect
During sentence comprehension semantic and world knowledge are rapidly integrated in parallel.
correct: The Dutch trains are yellow and very crowded. world knowledge violation: The Dutch trains are white and very crowded. semantic violation: The Dutch trains are sour and very crowded.
(300–550 ms) (300–550 ms)
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Gunter, Friederici & Schriefers, Journal of Cognitive Neuroscience, 2000
Amsterdam 29.08. 2014 >> Bilder >> Gunter
Phase 3: The integration phase: P600
Semantic and syntactic information interact during integration phase. The syntactic P600 is modulated by low cloze probability.
gender agreement gender disagreement
High cloze probability Low cloze probability
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Binder et al., Cerebral Cortex, 2009
Amsterdam 29.08. 2014 >> Bilder >> Gunter
+46 –46
The semantic network
The semantic network has been demonstrated to be of considerable size, involving inferior frontal and temporal regions in both hemispheres.
Left hemisphere Right hemisphere
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Friederici et al., Cerebral Cortex, 2006
3.09
left hemisphere
The syntactic network: Syntactic complexity
Functional subparts of the syntactic network are well localizable when well controlled stimulus material is used.
Broca's area Complexity-activation
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Zaccarella & Friederici, Poster presented at FENS, 2013
The syntactic network: Merge
Merge, the most basic syntactic operation of binding two elements together to form a phrase can be localized in a subpart of Broca's area, the ventral-anterior region C3.
Mass localization with cluster overlap
PH>LS activation Volume-of-interest
Amsterdam, 29.08.2014 >> Bilder >>Zaccarella
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Orange: main effect of hierarchy Light Blue: main effect of distance Red: interaction
2103_CNS_2009.ppt: 050309_neues Bild_100408 Fig 2_V3
Yellow: BA 45
Source: Makuuchi, Bahlmann, Anwander & Friederici, PNAS, 2009
Local syntactic network
Local syntactic network consist of BA 44 (syntactic hierarchy) and inferior frontal sulcus (syntactic WM). These regions interact during the processing of hierarchically structured sentences.
Processing embedded sentences
Green: BA
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Functional activation Structural connectivity
2511_bahlmann_fig3.fh11 2505_Leipzig_2006.ppt, umgefärbt: TractographyData.jpg
Source: Friederici, Bahlmann et al., PNAS, 2006
from BA44 to STG BA 44
Source: Friederici et al., NeuroReport, 2009
Long-range syntactic network
Long-range syntactic network consists of BA 44 (syntactic hierarchy), and pSTG/STS (integration).
BA44 pSTG/STS
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
29.08.2014 Amsterdam >>Bilder >> Meyer
Sources: Meyer, Grigutsch, Schmuck, Gaston & Friederici, Poster presented at CNS, 2014
The frontal and parietal/temporal region involved in syntactic processing synchronize (theta oscillations) during processing of syntactically complex sentences.
Neural synchronization within the syntactic network
Theta synchronization
Source localization Left frontal-parietal coherence
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Syntax (LH) – Prosody (RH) – Interaction (CC)
Source: Friederici, Kotz & von Cramon, Neuron, 2007
Controls
effect
Anterior CC lesion
effect
Posterior CC lesion
pros correct pros incorrect
Patients with lesions in the posterior Corpus Callosum do not show mismatch effect indicating that prosodic information (RH) cannot misguide the syntactic parser (LH).
–6 µV
0.8 s
PZ
6
–6 µV
0.8 s
PZ
6
–6 µV
0.8 s
PZ
6
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Sources: Zilles, Bacha-Trams, Palomero-Gallogher, Amunts & Friederici, Cerebral Cortex, 2015
29.08.2014 Amsterdam >>Bilder >> Bacha Trams
Areas that are part of the larger language network (red) may be neuroreceptorarchitectonically similar thereby providing a molecular basis for large-scale interaction.
The language network at the molecular level
Distribution of neuro-receptors Red: Language network
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Hierarchical cluster tree Multidimensional scaling
Sources: Zilles, Bacha-Trams, Palomero-Gallogher, Amunts & Friederici, Cerebral Cortex, 2015
29.08.2014 Amsterdam >>Bilder >> Bacha Trams
Areas that are part of the language network show a more similar multi-receptor organisation compared to other regions.
The language network at the molecular level
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Conclusion I
All the different levels of neurobiological analyses are relevant for our understanding of the brain-language relationship. Specific levels of description and their analytic methods should be applied for specific (linguistic) questions. But unless these level-specific analyses stay informed about the other levels, they will not advance science in the neurobiology of language.
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences
Conclusion II
Unless we – as those being interested in the brain-language relationship – do not learn to consider more than one level of neurobiological description, the scientific community will not recognize the importance of our work, which in the end is crucial for our understanding of what it means to be human.
Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences